Objective The present study investigated whether self-vocalization enhances auditory neural responsiveness to voice pitch feedback perturbation and how this vocalization-induced neural modulation can be affected by the extent of the feedback deviation. Method Event related potentials (ERPs) were recorded in 15 subjects in response to +100, +200 and +500 cents pitch-shifted voice auditory feedback during active vocalization and passive listening to the playback of the self-produced vocalizations. Result The amplitude of the evoked P1 (latency: 73.51 ms) and P2 (latency: 199.55 ms) ERP components in response to feedback perturbation were significantly larger during vocalization than listening. The difference between P2 peak amplitudes during vocalization vs. listening was shown to be significantly larger for +100 than +500 cents stimulus. Conclusion Results indicate that the human auditory cortex is more responsive to voice F0 feedback perturbations during vocalization than passive listening. Greater vocalization-induced enhancement of the auditory responsiveness to smaller feedback perturbations may imply that the audio-vocal system detects and corrects for errors in vocal production that closely match the expected vocal output. Significance Findings of this study support previous suggestions regarding the enhanced auditory sensitivity to feedback alterations during self-vocalization, which may serve the purpose of feedback-based monitoring of one’s voice.

BackgroundThe motor-driven predictions about expected sensory feedback (efference copies) have been proposed to play an important role in recognition of sensory consequences of self-produced motor actions. In the auditory system, this effect was suggested to result in suppression of sensory neural responses to self-produced voices that are predicted by the efference copies during vocal production in comparison with passive listening to the playback of the identical self-vocalizations. In the present study, event-related potentials (ERPs) were recorded in response to upward pitch shift stimuli (PSS) with five different magnitudes (0, +50, +100, +200 and +400 cents) at voice onset during active vocal production and passive listening to the playback.ResultsResults indicated that the suppression of the N1 component during vocal production was largest for unaltered voice feedback (PSS: 0 cents), became smaller as the magnitude of PSS increased to 200 cents, and was almost completely eliminated in response to 400 cents stimuli.ConclusionsFindings of the present study suggest that the brain utilizes the motor predictions (efference copies) to determine the source of incoming stimuli and maximally suppresses the auditory responses to unaltered feedback of self-vocalizations. The reduction of suppression for 50, 100 and 200 cents and its elimination for 400 cents pitch-shifted voice auditory feedback support the idea that motor-driven suppression of voice feedback leads to distinctly different sensory neural processing of self vs. non-self vocalizations. This characteristic may enable the audio-vocal system to more effectively detect and correct for unexpected errors in the feedback of self-produced voice pitch compared with externally-generated sounds.

Speaking is one of the most complex motor behaviors developed to facilitate human communication. The underlying neural mechanisms of speech involve sensory-motor interactions that incorporate feedback information for online monitoring and control of produced speech sounds. In the present study, we adopted an auditory feedback pitch perturbation paradigm and combined it with functional magnetic resonance imaging (fMRI) recordings in order to identify brain areas involved in speech production and motor control. Subjects underwent fMRI scanning while they produced a steady vowel sound /a/ (speaking) or listened to the playback of their own vowel production (playback). During each condition, the auditory feedback from vowel production was either normal (no perturbation) or perturbed by an upward (+600 cents) pitch shift stimulus randomly. Analysis of BOLD responses during speaking (with and without shift) vs. rest revealed activation of a complex network including bilateral superior temporal gyrus (STG), Heschl's gyrus, precentral gyrus, supplementary motor area (SMA), Rolandic operculum, postcentral gyrus and right inferior frontal gyrus (IFG). Performance correlation analysis showed that the subjects produced compensatory vocal responses that significantly correlated with BOLD response increases in bilateral STG and left precentral gyrus. However, during playback, the activation network was limited to cortical auditory areas including bilateral STG and Heschl's gyrus. Moreover, the contrast between speaking vs. playback highlighted a distinct functional network that included bilateral precentral gyrus, SMA, IFG, postcentral gyrus and insula. These findings suggest that speech motor control involves feedback error detection in sensory (e.g. auditory) cortices that subsequently activate motor-related areas for the adjustment of speech parameters during speaking.

The neural responses to sensory consequences of a self-produced motor act are suppressed compared with those in response to a similar but externally generated stimulus. Previous studies in the somatosensory and auditory systems have shown that the motor-induced suppression of the sensory mechanisms is sensitive to delays between the motor act and the onset of the stimulus. The present study investigated time-dependent neural processing of auditory feedback in response to self-produced vocalizations. ERPs were recorded in response to normal and pitch-shifted voice auditory feedback during active vocalization and passive listening to the playback of the same vocalizations. The pitch-shifted stimulus was delivered to the subjects’ auditory feedback after a randomly chosen time delay between the vocal onset and the stimulus presentation. Results showed that the neural responses to delayed feedback perturbations were significantly larger than those in response to the pitch-shifted stimulus occurring at vocal onset. Active vocalization was shown to enhance neural responsiveness to feedback alterations only for nonzero delays compared with passive listening to the playback. These findings indicated that the neural mechanisms of auditory feedback processing are sensitive to timing between the vocal motor commands and the incoming auditory feedback. Time-dependent neural processing of auditory feedback may be an important feature of the audio-vocal integration system that helps to improve the feedback-based monitoring and control of voice structure through vocal error detection and correction.

Objective The present study examined the differential effects of voice auditory feedback perturbation direction and magnitude on voice fundamental frequency (F0) responses and event-related potentials (ERPs) from EEG electrodes on the scalp. Methods The voice F0 responses and N1 and P2 components of ERPs were examined from twelve right-handed speakers when they sustained a vowel phonation and their mid-utterance voice pitch feedback was shifted ±100, ±200, and ± 500 cents with 200 ms duration. Results Downward voice pitch feedback perturbations led to larger voice F0 responses than upward perturbations. The amplitudes of N1 and P2 components were larger for downward compared with upward pitch shifts for 200 and 500 cents stimulus magnitudes. Shorter N1 and P2 latencies were also associated with larger magnitudes of pitch feedback perturbations. Conclusion Corresponding changes in vocal and neural responses to upward and downward voice pitch feedback perturbations suggest that the N1 and P2 components of ERPs reflect neural concomitants of the vocal responses. Significance The findings of interactive effects between the magnitude and direction of voice feedback pitch perturbation on N1 and P2 ERP components indicate that the neural mechanisms underlying error detection and correction in voice pitch auditory feedback are differentially sensitive to both the magnitude and direction of pitch perturbations.

The present study describes a technique for analysis of vocal responses to auditory feedback pitch perturbations in which individual trials are first sorted according to response direction and then separately averaged in groups of upward or downward responses. In experiment 1, the stimulus direction was predictable (all upward) but magnitude was randomized between þ100, þ200, or þ500 cents (unpredictable). Results showed that pitch-shift stimuli (PSS) of þ100 and þ200 cents elicited significantly larger opposing (compensatory) responses than þ500 cent stimuli, but no such effect was observed for "following" responses. In experiment 2, subjects were tested in three blocks of trials where for the first two, PSS magnitude and direction were predictable (block 1þ100 and block 2-100 cents), and in block 3, the magnitude was predictable (6100 cents) but direction was randomized (upward or downward). Results showed there were slightly more opposing than following responses for predictable PSS direction, but randomized directions led to significantly more opposing than following responses. Results suggest that predictability of stimulus direction and magnitude can modulate vocal responses to feedback pitch perturbations. The function and causes of the opposing and following responses are unknown, but there may be two different neural mechanisms involved in their production.

The neural mechanisms underlying processing of auditory feedback during self-vocalization are poorly understood. One technique used to study the role of auditory feedback involves shifting the pitch of the feedback that a speaker receives, known as pitch-shifted feedback. We utilized a pitch shift self-vocalization and playback paradigm to investigate the underlying neural mechanisms of audio-vocal interaction. High-resolution electrocorticography (ECoG) signals were recorded directly from auditory cortex of 10 human subjects while they vocalized and received brief downward (−100 cents) pitch perturbations in their voice auditory feedback (speaking task). ECoG was also recorded when subjects passively listened to playback of their own pitch-shifted vocalizations. Feedback pitch perturbations elicited average evoked potential (AEP) and event-related band power (ERBP) responses, primarily in the high gamma (70–150 Hz) range, in focal areas of non-primary auditory cortex on superior temporal gyrus (STG). The AEPs and high gamma responses were both modulated by speaking compared with playback in a subset of STG contacts. From these contacts, a majority showed significant enhancement of high gamma power and AEP responses during speaking while the remaining contacts showed attenuated response amplitudes. The speaking-induced enhancement effect suggests that engaging the vocal motor system can modulate auditory cortical processing of self-produced sounds in such a way as to increase neural sensitivity for feedback pitch error detection. It is likely that mechanisms such as efference copies may be involved in this process, and modulation of AEP and high gamma responses imply that such modulatory effects may affect different cortical generators within distinctive functional networks that drive voice production and control.

The present study investigated how pitch frequency, a perceptually relevant aspect of periodicity in natural human vocalizations, is encoded in Heschl's gyrus (HG), and how this information may be used to influence vocal pitch motor control. We recorded local field potentials from multicontact depth electrodes implanted in HG of 14 neurosurgical epilepsy patients as they vocalized vowel sounds and received brief (200 ms) pitch perturbations at 100 Cents in their auditory feedback. Event-related band power responses to vocalizations showed sustained frequency following responses that tracked voice fundamental frequency (F0) and were significantly enhanced in posteromedial HG during speaking compared with when subjects listened to the playback of their own voice. In addition to frequency following responses, a transient response component within the high gamma frequency band (75-150 Hz) was identified. When this response followed the onset of vocalization, the magnitude of the response was the same for the speaking and playback conditions. In contrast, when this response followed a pitch shift, its magnitude was significantly enhanced during speaking compared with playback. We also observed that, in anterolateral HG, the power of high gamma responses to pitch shifts correlated with the magnitude of compensatory vocal responses. These findings demonstrate a functional parcellation of HG with neural activity that encodes pitch in natural human voice, distinguishes between self-generated and passively heard vocalizations, detects discrepancies between the intended and heard vocalization, and contains information about the resulting behavioral vocal compensations in response to auditory feedback pitch perturbations.